Transactions  of  The  Academy  of  Science  of  St.  Louis. 

VOL,.  XI.  No.  7. 


THE  PROGRESS  MADE  IN  BOTANY  DURING  THE 
NINETEENTH  CENTURY. 

WILLIAM  TRELEASE. 


Issued  November  26 , 1901 . 


'§30*3 

Tit? 


hY 


o - 


^ o 


THE  PROGRESS  MADE  IN  BOTANY  DURING  THE 
NINETEENTH  CENTURY.* 

William  Trelease. 

WHAT  BOTANY  STOOD  FOR  AT  THE  BEGINNING  OF  THE  CENTURY. 

At  the  beginning  of  the  Nineteenth  Century  about  25,000 
species  of  plants  had  been  described,  and,  though  consider- 
| able  use  had  long  been  made  of  other  species  which  at  the 
beginning  of  the  century  were  unclassified  and  unnamed  by 
botanists,  the  number  of  these  were  relatively  small,  so  that 
the  entire  knowledge  of  botany,  economic  as  well  as  scientific, 
and  in  all  of  its  branches,  was  practically  confined  to  the 
limited  number  of  species  mentioned.  This  knowledge  con- 
sisted in  a recognition  of  their  specific  differences  and  the 
rather  superficial  affinities  and  relationship  deduced  from 
these  in  a great  but  often  hopelessly  scattered  and  frequently 
erroneous  literature,  and  in  popular  acquaintance  with  their 
useful  properties  — particularly  their  medicinal  virtues,  and  a 
general  blocking  out  of  their  anatomy  and  physiology, — to 
no  small  extent  a matter  of  subjective  opinion. 

SYSTEMATIC  BOTANY. 

About  the  middle  of  the  preceding  century,  Linnaeus  had 
elaborated  a workable,  if  artificial,  system  of  classification, 
which,  with  brief  but  sharp  diagnoses,  made  it  reasonably  easy 
to  ascertain  whether  a given  species  of  plant  in  hand  had  been 
previously  described  or  was  new  to  science;  and  as  he  had 
combined  with  this  the  very  simple  expedient  of  referring  to 
the  several  species  by  latinized  binomials  instead  of  by  de- 
scriptive phrases,  the  naming  and  describing  of  species  has 
proved  not  only  one  of  the  most  necessary  but  also  one  of  the 
easiest  and  most  popular  branches  of  this  as  well  as  of  the 
related  biologic  science,  zoology,  during  the  century  just 
closed. 


* An  address  delivered  before  The  Academy  of  Science  of  St.  Louis, 
November  8,  1901. 


(125) 


126 


V 


Trans.  Acad.  Sci.  of  St.  Louis. 


Linnaeus  himself,  in  1771  admitted  sssi 
of  which  more  than  nine-tenths  beSol  to  ZT  f f “tS 
and  grossly  marked  o-r0up  the  Aowp  •°  h st  obvri0Uf 
the  ferns,  still  represent  the  field  of  Zfnytr 
persons.  The  rapidity  of  progress  in  diff  • Unec*ucated 

nized  species  and  characterizing  such  af  had  r ? 

served  is  shown  by  the  increase  of  I inn  . ma‘ned  unob- 
some  70,000  before  the  fiZT  f Linnaeus  s scant  9,000  to 

studies  of  Presl,  Milde  the  * ’ , ,h  throuSh  the 

:?*7r  ~ » tbe  „„z. 

oi  *.  rzpxz  r:i 
F ri±~ 

ongei  believed  to  be  necessary  for  the  flowering  plants  of  the 
more  accessible  parts  of  the  world,  these  having  been  foil 
satisfactorily  disposed  of  on  the  grosser  or  so  called  T • 7 

ideas  of  specific  limitation,  and  partly  because  of  D“iw! 
tudies  of  parasitism  and  a recognition  that  many  of  the 
diseases  of  cultivated  plants  are  caused  by  fungi  the  diff 

of  which  been,,  in,p„rt„,/f„  eco"“ [- 

as  well  as  a systematic  point  of  view  At  the  ol -pTu 
century  not  far  from  180,000  species  of  plants  were  known' 
of  which  some  75,000,  or  more  than  the  total  number  of 
species  known  ,n  all  groups  at  the  end  of  the  first  quarter  of 
the  century,  are  cryptogams.  The  last  decade,  however  has 
witnessed  a proportionally  greater  increase  in  phanerogam" 


Trelease — Botany  During  the  19th  Century.  127 

species  than  that  marking  the  immediately  preceding  decades, 
because  of  the  geographic  exploration  already  referred  to, 
and  still  more  because  of  a growing  change  in  the  scale  of 
specific  differentials  which  has  resulted  in  the  segregation  of 
[many  forms  which  under  the  older  views  passed  for  at  the 
most  varieties  of  polymorphic  or  variable  species.  The 
genera  Ruhus  and  Hieracium,  in  Europe,  and  Viola , Sisyrin- 
chium  and  Crataegus , in  our  own  country,  well  illustrate  my 
meaning. 

Just  as  the  descriptive  manuals  of  Linnaeus,  and  the  editions 
of  them  published  after  his  death  by  Schultz,  Willdenow  and 
others,  facilitated  and  stimulated  the  accumulation  of  hith- 
erto unrecognized  species  at  the  beginning  of  the  century, 
its  progress  throughout  has  been  recorded  and  accelerated  by 
the  publication  of  later  works  of  the  same  general  character 
and  purpose.  For  the  flowering  plants,  some  of  the  most 
noteworthy  of  such  general  descriptive  works  are  the  incom- 
plete Prodromus  and  Monographiae  of  the  De  Candolles,  the 
numerous  revisions  of  genera  and  families  in  Engler’s  Jahr- 
biicher  and  the  Journal  of  the  Linnean  Society,  and  the  com- 
prehensive Index  Kewensis  prepared  by  Mr.  Jackson  under  a 
provision  made  in  Darwin’s  will ; and  no  account  of  this 
aspect  of  the  science  would  be  at  all  complete  without  refer- 
ence to  the  books  and  journals  devoted  to  the  illustration  of 
plants,  foremost  among  which  stands  the  Botanical  Maga- 
zine, which,  founded  by  Curtis  in  1790,  has  been  continued 
without  interruption,  and  at  the  end  of  1900  contained  7751 
colored  plates,  mostly  illustrative  of  plants  of  decorative 
value.  For  Pteridophytes,  the  manuals  of  Hooker  and  Baker 
have  been  most  helpful.  Bride],  Schimper  and  Warnstorf 
stand  out  prominent  among  those  who  have  published  compre- 
hensive manuals  of  the  Bryophytes,  while  the  enormous 
Sylloge  Fungorum  of  Saccardo  and  the  as  yet  incomplete 
Sylloge  Algarum  of  DeToni  have  made  accessible  the  myriads 
of  scattered  descriptions  of  species  belonging  to  these  groups 
of  the  lower  cryptogams. 

SYSTEMS  OF  CLASSIFICATION. 

The  simplicity  of  Linnaeus’s  classification  of  flowering 
plants  has  been  mentioned.  The  popular  handbooks  even  of 


128 


Trans.  Acad.  Sci.  of  St.  Louis. 


our  own  flora,  up  to  a point  somewhat  after  the  middle  of  thfe 
century,  were  based  on  this  system,  which,  when  the  purpose 
of  the  student  was  to  find  the  name  of  a plant,  has  scarcely 
been  equaled  by  any  other ; yet  it  had  one  very  great  defect,, 
in  that  plants  which  were  obviously  related  might  come  tc 
stand  far  apart  in  it,  so  that  the  suggestion  of  this  relation^ 
ship  would  be  lost  on  the  user  of  a book  in  which  it  was 
followed.  Even  before  the  close  of  the  preceding  century, 
efforts  had  been  turned  to  the  arrangement  of  a natural  se- 
quence of  the  higher  groups  of  plants,  so  that  those  which 
possess  a number  of  important  and  correlated  characters  in 
common  might  be  brought  together,  leaving  the  tracing  of 
any  given  species  to  its  place  in  the  system  for  a quite  inde- 
pendent artificial  key,  — the  Linnean,  for  instance,  or  some 
other  specially  fancied  by  the  writer  or  suited  to  his  purposes. 
To  the  Jussieus  the  inception  of  this  movement  in  a modern 
sense  is  due,  and  the  elder  DeCandolle  stands  out  prominently 
among  those  who  amplified  and  bettered  it;  and  yet  the  suc- 
cess of  these  earlier  seekers  for  a natural  system  was  but  par- 
tial, and  in  the  summation  of  their  conclusions,  as  exemplified, 
for  instance,  in  the  great  Genera  Plantarum  of  Bentham  and 
Sir  Joseph  Hooker,  though  many  of  the  resultant  groups, 
even  no  higher  than  orders,  possess  a very  puzzling  com- 
plexity because  of  the  insertion  of  aberrants,  there  still  re- 
main many,  as,  for  instance,  a large  part  of  those  constitut- 
ing the  so-called  Apetalae,  which  are  obviously  little  more 
than  makeshifts,  loose-jointed  in  themselves  and  with  scarce 
concealed  affiliations  of  the  most  diverse  kinds.  As  early  as 
the  middle  of  the  century,  by  his  comparative  developmental 
studies  of  the  gymnosperms  and  higher  cryptogams,  Hof- 
meister  laid  the  foundations  of  a more  rational  system,  which, 
largely  through  the  labors  of  Alexander  Braun  and  Eichler, 
culminated  in  the  phylogenetic  system  of  Engler,  which  marks 
the  close  of  the  century. 

Somewhat  comparable  needs  and  advances  have  marked  the 
knowledge  of  the  cryptogams.  The  ferns  and  their  allies 
early  differentiated  themselves  from  the  remainder  of  this 
great  second  group  of  Linnaeus,  and  the  mosses  and  liver- 
worts as  quickly  came  to  be  recognized  as  forming  another 


Trelease  — Botany  During  the  19th  Century.  129 

distinct  group  of  primary  importance  in  any  natural  classifica- 
tion, the  researches  of  Ho fmeister  contributing  largely  to  this 
result;  but  even  to-day,  convenience  of  treatment,  if  no  other 
reason,  causes  specialists  to  write  commonly  on  either  algae, 
fungi  or  lichens,  according  to  the  group  of  thallophytes  they 
may  be  studying.  And  yet  the  beginning  of  better  things 
has  been  made,  for  DeBary’s  suggestion  and  Schwendener’s 
morphological  demonstration  that  lichens  are  in  reality  only 
certain  fungi  with  enslaved  or  commensal  algae  as  an  integral 
and  usually  necessary  part  of  their  organization  marks  the 
close  of  the  third  quarter  of  the  century,  and  in  the  conclud- 
ing quarter  various  efforts  have  been  made  at  a classification 
of  the  thallophytes  on  more  scientific  grounds  than  the  pres- 
sence  or  absence  in  them  of  chromophyll-bearing  cells  or  tis- 
sues. Though  the  goal  may  not  yet  have  been  reached,  these 
efforts  are  full  of  promise  for  success  in  the  century  that  is 
now  on  the  calendar. 

EVOLUTION  AND  CLASSIFICATION. 

It  was  in  the  first  decade  of  the  century  that  Lamarck,  fol- 
lowing a line  of  thought  that  had  caused  men  long  before  his 
time  to  speculate  on  the  varied  forms  of  nature,  attempted 
to  show  how  environment,  use  and  disuse  of  parts,  and  sim- 
ilar natural  factors  might  have  brought  about  modifications 
leading  to  the  origin  of  new  species  from  ancestors  otherwise 
characterized;  and  the  year  1858  will  always  stand  out  in 
prominence  in  the  history  of  biology  because  of  the  simulta- 
neous presentation  in  that  year  of  almost  identical  explana- 
tions of  the  manner  in  which  natural  selection,  or  the  survival 
of  the  fittest  in  life’s  struggle,  might  and  of  necessity  must 
lead  to  the  repeopling  of  a given  territory  by  new  forms  de- 
scended from  those  pre-existing,  provided,  in  the  progress  of 
time,  the  conditions  of  life  were  changeful  and  variations 
were  present  in  offspring,  as  compared  with  one  another  and 
their  parents,  — as  was  well  known  to  be  the  case.  Darwin 
and  Wallace,  the  authors  of  these  first  papers,  did  not  go  to 
the  bottom  of  their  great  subject,  and  the  last  word  on  it  is 
far  from  having  been  said  yet,  but  the  theory  of  organic 
evolution  may  be  regarded  to-day  as  an  axiom  on  which  most 
philosophical  analyses  of  biology  rest  as  a footing  course. 


130 


Trans.  Acad.  Sci.  of  St.  Louis. 


Closely  connected  with  the  changing  conceptions  as  to  the^ 
origin  and  fixity  of  species,  was  a much  increased  interest  in 
such  evidence  concerning  the  plants  of  the  past  as  was  afforded 
by  their  fossil  remains,  and,  largely  through  the  work  of 
Brongniart,  Goeppert,  Heer,  the  elder  Schimper,  von  Ettings- 
hausen,  Saporta  and  Solms  Laubach,  and  Dawson,  New- 
berry, and  Lesquereux  in  this  country,  paleobotany  has 
assumed,  in  the  last  fifty  years,  a position  of  no  small  im- 
portance. 

Partly  because  of  the  same  reasons,  the  geographical  dis- 
tribution of  plants  and  the  influences  controlling  widespread 
or  restricted  occurrence  in  the  case  of  individual  genera  or 
species  has  also  assumed  an  importance  in  recent  years  not 
formerly  recognized  for  it,  and  on  the  foundation  laid  by 
DeCandolle,  Humboldt  and  Martius,  Grisebach,  Engler,  Drude 
and  the  younger  Schimper  have  grounded  a line  of  botanical 
research  in  which  morphologists,  systematists  and  evolution- 
ists are  alike  interested. 

With  the  change  in  the  world’s  view  of  the  fixity  of  species, 
and  of  their  several  and  independent  origin  in  their  present 
form,  came  new  and  somewhat  differently  conceived  efforts 
to  group  plants  in  a natural  system,  the  ultimate  object  being 
virtually  the  production  of  a classification  which  should  rep- 
resent descent  relationship  as  well  as  organic  or  morphological 
affinity,  and  which,  in  a word,  should  present  the  family  tree 
of  any  individual  group  or  species,  — to  the  primitive  animal 
and  vegetable  main  divisions  of  which  Haeckel  in  particular 
has  given  attention.  A comparative  glance  at  the  Genera 
Plantarum  of  Bentham  and  Hooker,  the  synopses  of  Van 
Tieghem  and  Warming,  and  the  still  incomplete  Pflanzen- 
familien  of  Engler  and  Prantl  will  show  how  great  have  been 
the  changes  wrought  in  systems  of  classification  by  the 
introduction  of  these  later  considerations  and  motives.  Free 
to  read  heredity  and  atavism  into  the  explanation  of  aberrant 
minor  characters,  rudiments  and  vestiges,  these  men  have  often 
found  in  the  minuter  details  of  anatomy,  reproduction  and 
development  most  surprising  indications  of  affinity  between 
superficially  and  externally  dissimilar  groups.  That  they  are 
not  at  one  in  their  conclusions,  indicates  that  the  Twentieth 


Trelease  — Botany  During  the  19th  Century.  131 

Century  may  regard  the  preparation  of  a truly  natural  system 
even  of  the  higher  plants  as  a part  of  its  legitimate  and 
necessary  work,  and  it  may  well  be  that  even  though  this  task 
be  accomplished,  a like  result  among  the  lower  cryptogams 
will  be  reserved  for  the  next  century.  At  any  rate,  although 
DeBary  and  others  have  contributed  to  a rational  comparison 
of  the  larger  groups  of  thallophytes,  a glance  at  the  sytematic 
memoirs  relating  to  the  fungi  and  algae  shows  a most  obvious 
if  convenient  artificialty  in  their  classification. 

MORPHOLOGY  AND  ANATOMY. 

Some  years  since,  I saw  with  much  interest  a palm  in  the 
Botanical  Garden  of  Padua  on  which,  toward  the  end  of  the 
Eighteenth  Century,  the  great  poet  Goethe  made  some  of  the 
observations  which  led  to  a formulation  of  his  theory  of  meta- 
morphosis in  the  parts  of  plants, — a theory  which,  in  the  first 
half  of  the  century  just  closed,  DeCandolle,  our  own  Engel- 
mann  and  others  put  upon  a more  scientific  basis  as  a funda- 
mental idea  in  plant  morphology.  Toward  the  middle  of  the 
century,  the  superficial  indications  afforded  by  position,  grada- 
tion and  malformation  of  parts  were  much  strengthened  by 
embryological  and  developmental  studies,  and  it  was  about  this 
time  that  the  details  of  cellular  structure,  grossly  known  for  a 
couple  of  centuries,  were  brought  out  by  Robert  Brown  and 
Schleiden,  the  latter  of  whom  stated  in  another  form  for 
plants  the  general  fact  of  the  origin  of  every  cell  from  a pre- 
vious cell,  succinctly  expressed  by  the  now  venerable  Virchow, 
whose  eightieth  birthday  has  recently  been  celebrated  in  this 
country  as  well  as  in  his  native  land ; for  by  this  time  these 
structures  had  come  to  be  recognized  as  the  seat  of  vital 
manifestations  through  their  protoplasm,  which,  discovered 
and  named  by  von  Mohl,  and  the  nuclear  differentiation  of 
which  was  observed  by  Robert  Brown  in  1835,  and  which  was 
shown  to  be  similar  in  animals  and  plants  by  Cohn  in  1850, 
Huxley  has  so  happily  designated  as  the  physical  basis  of  life. 

Though  external  morphology  and  anatomy,  the  latter  even 
in  some  of  its  minuter  details,  had  come  down  from  the  past, 
both  may  be  said  to  have  been  made  a part  of  science  in  the 
Nineteenth  Century,  and  the  fact  that  homologous  members 


132 


Trans.  Acad.  Sci.  of  St.  Louis. 


may  serve  the  most  diverse  organic  purpose,  that  sometimes 
analagous  organs,  like  the  leaf  of  the  moss  and  that  of  the 
flowering  plant,  cannot  be  morphologically  compared,  since 
they  are  parts  of  fundamentally  unlike  plant  bodies,  shown 
primarily  by  Hofmeister’s  discovery  of  alternating  genera- 
tions  in  1851  (one  representing  the  gametophyte  and  the 
other  the  sporophyte  of  beings  with  alternating  sexual  and 
non-sexual  generations),  and  that  cells,  cellular  tissues,  and 
systems  of  such  tissues  show  a similar  and  comparable  plia- 
bility in  their  adaptation  to  physiological  function,  as  Haber- 
land  and  others  have  made  clearly  evident,  with  many  other 
facts  of  equal  importance  for  a right  understanding  of  nature, 
may  be  credited  in  large  part  to  the  last  half,  and,  as  to  much 
of  their  detail,  to  the  last  quarter,  of  the  century.  Indeed, 
the  consideration  of  tissues  from  a proper  morphological 
point  of  view  dates  practically  from  Hanstein’s  studies  in 
1868,  and  their  rational  terminology  was  established  by 
DeBary  nearly  a decade  later. 

Though  initially  wrong,  Schleiden  as  early  as  1837  laid  the 
foundation  of  embryology  in  botany,  and  the  organogenetic 
studies  of  Hofmeister,  Payer,  Sachs  and  Goebel  will  always 
stand  as  classics  in  the  application  of  the  developmental  line 
of  research  to  the  progressively  formed  grosser  parts  of  more 
mature  plants. 

PHYSIOLOGY. 

Physiology,  either  of  animals  or  plants,  could  scarcely  have 
become  a science  before  the  determination  of  the  grosser 
chemical  composition  of  the  atmosphere,  which,  made  by  the 
chemist  Priestley  toward  the  end  of  the  Eighteenth  Century, 
was  quickly  followed  up  by  him,  Ingen-Housz,  de  Saussure, 
Hales  and  numerous  others,  with  the  result  of  showing  that 
a very  considerable  part  of  the  organic  matter  of  which 
plants  consist  is  derived  from  the  carbon  dioxide  of  the  at- 
mosphere, which  is  fixed  in  carbohydrate  form  in  the  green 
parts  of  plants  under  the  influence  of  light ; and  the  studies 
of  Draper  and  Wilhelm  Engelmann  stand  out  in  prominence 
as  contributing  to  our  present  knowledge  that  certain  wave- 
lengths of  sunlight,  when  passing  through  the  chlorophyll 
or  comparable  pigments  of  plants,  disappear  as  light,  and  are 


Trelease  — Botany  During  the  19th  Century.  133 

converted  into  chemical  or  physical  energy,  which,  under 
the  guidance  of  the  living  protoplasm  of  the  cells,  is  utilized 
for  the  breaking  down  of  carbon  dioxide  and  water,  their 
elements  being  then  recombined  into  the  organic  products 
referred  to,  the  most  usually  recognizable  of  which  is  starch. 
An  attendant  liberation  of  oxygen,  constituting,  with  the  ab- 
straction of  carbon  dioxide,  a purification  of  the  air,  so  far 
as  the  needs  of  animals  are  concerned,  was  made  known 
shortly  before  the  century  began,  but  it  is  to  Saussure,  at  its 
very  beginning,  that  the  connection  of  this  with  actual  plant 
nutrition  is  due,  and  it  was  he,  too,  who  gave  the  first 
clear  demonstration  that  the  remainder  of  plant  food  is  de- 
rived from  the  soil.  A detailed  study  of  this  subject,  as 
well  as  of  the  metabolism  or  elaboration  and  transmutation  that 
food  undergoes  in  the  plant  in  its  various  nutritive  and  storage 
processes,  occupied  particularly  Sachs  during  the  third  quarter, 
and  Pfeifer  during  the  last  quarter  of  the  century,  Pfeifer’s 
ingenious  investigation  of  the  osmotic  action  of  root  hairs  being 
particularly  interesting  in  connection  with  the  physical  prob- 
lems of  the  absorption  of  crude  materials  and  the  retention  of 
organic  products  in  the  self-same  organ.  The  last  half  of  the 
century  has  also  produced  the  demonstration,  on  a large  scale 
in  the  field  experiments  of  Gilbert  and  Lawes,and  on  a smaller 
scale,  but  under  more  rigid  control,  in  the  laboratories  of  nu- 
merous investigators,  of  the  fact  that  while  free  atmospheric 
nitrogen  is  not  available  for  the  nutrition  of  higher  plants* 
which  therefore  as  a rule  require  for  their  proper  support  an 
abundance  of  available  nitrogen  supplied  to  the  roots  in  the 
form  of  nitrates,  nitrites,  etc.,  the  Leguminosae  as  a class 
make  use  of  large  quantities  of  this  atmospheric  nitrogen, 
not,  indeed,  in  its  free  form  directly,  but  through  the  inter- 
vention of  certain  of  the  lowest  fungi  which  inhabit  their 
roots  as  parasites,  but,  having  the  power  of  assimilating 
nitrogen  in  forms  in  which  it  is  not  usable  by  the  higher 
plants,  contribute  to  the  latter  enough  of  the  product  of  their 
own  activity  to  more  than  compensate  for  whatever  injury 
they  may  cause  by  their  parasitic  invasion  of  the  tissues  of 
the  host.  Indeed,  pure  cultures  of  these  pseudo-parasites 
are  on  the  market,  under  the  name  of  nitragin,  for  the  inoc- 


134 


Trans.  Acad.  Sci.  of  St.  Louis. 


ulation  of  new  soil  when  sown  to  clover  and  other  legumin- 
ous crops,  though  it  must  be  added  that  the  practical  value 
of  this  inoculation  is  thrown  in  considerable  doubt  by  re- 
cently made  laboratory  experimental  tests. 

PROTOPLASM. 

Doubtless  the  most  important  of  all  discoveries  in  physiol- 
ogy is  that  of  protoplasm  as  the  living  working  part  of  both 
plants  and  animals,  in  the  early  phases  of  which  von  Mohl, 
Robert  Brown,  Naegeli  and  Cohn  played  a prominent  part. 
Studies  on  this  substance,  its  physical  and  chemical  properties, 
and  its  activity,  have  occupied  many  of  the  best  chemical, 
physical  and  biological  investigators  of  the  last  half  of  the 
century,  and  are  destined  to  be  the  keystone  of  physiological 
attainments  in  the  century  we  are  now  entering  upon. 

Though  sex  in  the  flowering  plants  was  known  long  before 
the  century  opened,  to  the  extent  that  the  co-operation  of 
stamen  and  pistil,  and  even  the  transfer  of  pollen  from  the 
former  to  the  latter,  was  recognized  as  necessary  for  the  pro- 
duction of  fertile  seed, — a fact,  indeed,  which  Linnaeus  in- 
dicated and  even  amplified  in  his  designation  of  the  groups 
which  he  called  phanerogams  and  cryptogams, — it  was  not 
until  1823  that  Amici  observed  the  growth  of  the  pollen  tube 
to  the  ovule,  and  real  fertilization,  the  union  of  protoplasmic 
structures,  was  not  demonstrated  until  the  close  of  another 
quarter  of  a century,  when  Hofmeister  and  Pringsheim  at  in- 
tervals of  a few  years  described  it  respectively  for  some  of 
the  higher  and  lower  cryptogams. 

The  greatest  advance  in  protoplasmic  study  was  doubtless 
made  possible  by  Strasburger’s  introduction,  in  1875,  of 
methods  for  fixing  protoplasmic  structures  in  certain  desired 
states  of  their  transformations,  by  the  use  of  killing  and 
hardening  fluids,  and  the  addition  a few  years  later  of  dif- 
ferential staining  processes,  as  a result  of  which,  largely 
through  his  efforts  and  those  of  his  pupils,  the  minutiae  of 
both  cell  division  and  cell  union  have  been  carried  to  a won- 
derful detail, — perhaps  the  least  expected  result  of  which  is 
the  closing  discovery  of  the  century  of  an  unexplained  double 
fertilization  in  the  case  of  the  flowering  plants,  by  which  the 
endosperm  is  formed  as  well  as  the  embryo. 


Trelease  — Botany  During  the  19th  Century.  135 

How  protoplasm  carries  “ life,”  the  nature  of  the  reactions 
Vt  shows  to  stimuli  of  various  kinds,  causing  it  to  work,  to 
change,  to  rest,  to  die,  how  it  is  moved  to  vary  in  the  forms 
of  tissues  and  organs  over  the  construction  of  which  it  pre- 
sides, how  it  transmits  characters  of  form  and  action  from 
parent  to  offspring  and  reverts  now  and  then  to  ancestral 
structures  and  traits  in  both  animals  and  plants,  are  scarce 
more  than  question  marks  on  an  otherwise  clean  page  spread 
out  before  the  Twentieth  Century,  and  it  is  not  possible  yet 
to  say  whether  they  will  receive  their  answer  soon  or  always 
remain  unanswered. 

ecology. 

One  of  the  most  popular  lines  of  physiological  work  to-day 
concerns  itself  with  special  modifications  and  activities  con- 
nected with  local  environment  and  what  may  be  called  the 
personal  or  individual  needs  of  plants,  in  contrast  with  their 
needs  as  a class.  This  is  called  biology  by  some  and  ecology 
by  others. 

Just  before  the  end  of  the  Eighteenth  Century,  a German, 
Sprengel,  observed  a few  hairs  springing  from  the  base  of  the 
petals  of  a wild  geranium,  and,  though  he  did  not  share  the 
impersonal  teleological  views  that  prevail  to-day,  he  believed 
that  these  hairs  existed  for  a purpose,  which  he  undertook  to 
find  out.  Under  them  he  found  glands  secreting  a sweet 
fluid,  nectar,  which  he  saw  was  sheltered  by  them,  but  the 
nectar  was  a further  puzzle.  Bees  came  to  the  flowers  as  he 
watched,  and  removed  the  nectar,  which  the  glands  had 
secreted  and  the  hairs  protected  for  them,  and  the  question 
seemed  answered ; for  an  idea,  somewhat  prevalent  even  yet, 
that  everything  exists  for  the  good  of  something  else,  — gen- 
erally higher  in  the  scale  than  itself, — was  commonly  held 
in  his  day.  Further  observation,  however,  showed  him  that 
the  bees  became  dusted  with  pollen  and  that  they  uncon- 
sciously transferred  some  of  this  to  the  stigmas  of  the  flowers, 
while  rifling  them  of  their  sweets,  and  that  this  transfer, 
long  known  as  necessary  in  some  manner  for  fertilization  and 
the  quickening  of  the  germ,  could  not  otherwise  take  place 
except  by  remote  chance.  Then  he  examined  many  other 
kinds  of  flowers,  and  reached  the  broad  conclusion  that  nectar 


136 


Trans.  Acad.  Sci.  of  St.  Louis. 


in  these  organs  exists  for  the  sole  purpose  of  attracting  to 
them  insects,  sometimes  of  one,  sometimes  of  another  kind 
(for  which  it  is  protected  from  rain  and  dew  and  commonly 
from  other  classes  of  insects,  and  to  which  its  presence  is 
made  known  by  odor  and  color,  and  its  position  by  grooves 
and  other  guiding  mechanism  and  by  variegation  in  the  col- 
oring), which,  while  serving  their  own  purposes,  ensure  the 
pollination  of  some  flowers  which  might  attain  the  same  end 
directly  as  well  of  others  which  from  some  seeming  freak  of 
nature  mature  stamens  and  pistils  at  different  times  or  even 
have  them  separated  in  different  flowers,  — sometimes,  even, 
on  different  individuals.  A half  century  later,  Mr.  Darwin, 
seeing  in  floral  forms,  colors  and  odors  something  more  than 
means  of  overcoming  chance  defects  in  plan  or  development, 
showed  not  only  the  general  accuracy  of  Sprengel’s  conclu- 
sions as  illustrated  by  a host  of  other  cases,  but  that  they 
might  be  carried  a step  further,  by  stating  the  purpose  of  the 
structural  and  functional  peculiarities  in  question  to  be  the 
effecting  of  cross  fertilization.  Then  he  set  to  work  to  prove, 
by  a long-continued  series  of  experiments,  whether  or  not 
this  is  connected  with  a gain  to  the  offspring  resulting  from 
such  crosses,  and  we  cannot  question  the  resulting  conclusion 
that  it  is.  Indeed  it  may  be  asked  if  any  axiom  is  more  im- 
portant to  an  understanding  of  the  evolutionary  adaptation  of 
species  to  changing  environment  than  the  obvious  conclusion 
that  sex,  and  particularly  the  partition  of  the  sexes  with  sec- 
ondary provisions  of  the  most  varied  kinds  for  their  functional 
union,  is  a most  potent  factor  for  the  introduction  of  variation 
within  helpful  limits,  on  which  natural  selection  may  build 
with  the  current  of  the  times,  as  well  as  for  the  direct  bet- 
terment of  the  offspring. 

How  dissemination  is  effected,  and  the  structures  connected 
with  it;  how  plants  may  climb  to  the  light  and  air  with  a 
minimum  expenditure  of  material,  over  their  more  robust 
competitors  when  the  latter  have  reached  their  own  limit  in 
the  occupation  of  the  soil ; how  they  may  feed  upon  each 
other  and  upon  animals  ; how  they  may  extend  into  deserts 
and  the  salt  sea:  — these  and  many  other  questions  show  the 
range  of  ecology  as  it  is  now  occupying  alike  physiologists. 


Trelease  — Botany  Daring  the  19th  Century.  137 

morphologists  and  systematists,  and,  while  much  remains  to 
be  done,  its  blocking  out  is  likely  to  stand  as  one  of  the  more 
important  achievements  of  the  century  just  closed. 

APPLIED  BOTANY. 

Hand  in  hand  with  the  advance  of  pure  botany,  and  largely 
dependent  upon  it,  have  gone  at  least  as  great  advances  in  the 
application  of  ascertained  facts;  and  the  best  agricultural 
practice  of  to-day,  as  exemplified  in  the  intelligent  use  of 
fertilizers,  the  rotation  of  crops,  etc.,  is  conformed  to  the 
teachings  of  vegetable  physiology,  while  the  knowledge  of 
the  plasticity  of  plants  has  made  each  of  the  later  decades 
the  recipient  of  numerous  improved  races  and  varieties  of 
cultivated  species.  To-day,  among  the  more  pliable  forms, 
within  certain  limits  that  cannot  yet  be  overstepped,  new 
varieties  suited  to  special  needs  are  selected  and  bred  by  men 
like  Burbank  with  surprising  rapidity  and  accuracy,  almost 
to  drawing  and  specification,  because  of  the  practical  appli- 
cation of  the  knowledge  that  plants  are  plastic  under  environ- 
ment and  selection. 

The  details  of  other  contributions  of  botanical  science  to 
human  needs  are  of  no  less  interest.  Modern  brewing  is 
carried  on  scientifically,  as  a result  of  the  fermentation  studies 
of  Schwann  and  Pasteur  and  the  cultural  investigations  of 
Hansen,  a yeast  being  employed  which  has  developed  from 
a single  cell  of  known  pedigree  and  properties.  Citric  acid 
and  vinegar  are  produced  with  equal  certainty  if  less  com- 
plexity of  manipulation,  and  the  method  of  pure  cultures 
of  the  necessary  ferments  is  coming  into  considerable  use  in 
the  ripening  of  cream  for  butter  and  of  cheese. 

Perhaps  the  most  markedly  useful  application  of  the  botan- 
ical knowledge  of  the  century  is  in  the  field  of  medicine.  In 
the  early  part  of  the  century,  the  physician  was  of  necessity 
a botanist,  and  indeed  many  of  the  botanists  whose  names  ap- 
pear in  this  account  were  physicians  by  training.  From  the 
Middle  Ages  he  had  the  knowledge  of  physic  that  character- 
izes primitive  man  everywhere  to-day,  and  this  had  gradually 
come  to  represent  a pseudo-science  of  therapy  which  he  prac- 
ticed by  diagnosis,  prescription  and  exhibition, — if  I may 


138 


Trans.  Acad.  Sci.  of  St.  Louis. 


borrow  a word.  But  the  century  just  closed  has  seen  a dif- 
ferentiation of  pharmacy  from  medicine  which  has  not  only 
greatly  simplified  the  materia  medica  through  its  more 
careful  investigation,  but  has  given  the  physician  more  free- 
dom to  follow  out  his  own  field,  so  that  to-day,  while  he  must 
know  experimentally  the  physiological  action  of  more  plants 
than  his  predecessors  actually  used,  he  need  not  ordinarily 
know  more  of  these  plants  than  that  their  active  principles,  in 
sulphates,  fluid  extracts  and  the  like  are  commercially  pro- 
curable in  definite  degrees  of  assimilability  and  concentration, 
though  his  final  trials  have  not  been  lessened  thereby. 

The  century  will  forever  stand  as  that  in  the  last  third  of 
which  the  germ  causation  of  disease  was  made  known,  and 
the  names  of  Pasteur,  Koch  and  Lister  are  inseparably  con- 
nected with  this  great  addition  to  knowledge,  which, — since 
the  germs  of  disease  are  for  the  most  part  bacteria,  that, 
though  of  simple  and  aberrant  structure,  are  commonly 
classed  with  plants,  — must  be  counted  among  the  achieve- 
ments of  botany.  Sanitation  and  surgery  have  both  been  put 
on  an  entirely  new  footing  by  this  recognition  that  the  minut- 
est organisms  yet  known  are  responsible  for  many  of  the  most 
dreaded  pests,  so  that  the  exclusion  or  elimination  of  germs, 
and  the  use  of  their  own  products, — either  direct  or  by  ani- 
mal reaction  in  the  form  of  serums, — in  therapy,  form  to- 
day the  surest  safeguard  against  infectious  disease,  the  occur- 
rence of  which  may  soon  be  regarded  as  almost  a stigma  on 
civilization. 

The  century  just  closed  has  witnessed  an  almost  equal  ad- 
vance in  knowledge  of  the  causation  of  the  diseases  of  plants 
themselves.  Rusts,  smuts  and  mildews  are  no  longer  looked 
upon  as  exanthemata,  but  the  fruits  of  parasitic  fungi,  which, 
more  than  is  the  case  with  the  parasites  of  animals,  are  of  the 
less  minute  and  therefore  more  easily  seen  and  controlled 
groups, — though  plants  are  also  subject  to  a few  bacterial 
diseases.  Much  has  been  done  in  the  way  of  prophylaxis, 
and  something  in  the  way  of  germicide  therapy,  in  this  field, 
and  the  foundations  of  a true  science  of  plant  pathology  based 
upon  distorted  physiological  processes  due  to  improper  en- 
vironment, food  and  the  like,  or  to  the  ferments  secreted  by 


Trelease — Botany  During  the  19tli  Century.  139 

parasites  or  the  chemical  alterations  which  these  induce  in  the 
affected  plants,  may  be  said  to  have  been  laid  in  the  closing 
days  of  the  century  by  Professor  Marshall  Ward. 

POPULARIZATION  AND  PUBLICATION. 

The  development  of  any  department  of  science  is  closely 
connected  with  its  power  of  interesting  men.  The  present 
tendency  of  this  interest  is  more  and  more  commercial  and 
economic,  though  it  should  be  said  at  the  same  time  that  no 
earlier  period  has  witnessed  a higher  development  of  interest 
in  the  purely  abstract  problems  of  science. 

The  lucid,  terse  Latin  of  Linnaeus  did  much  to  popularize 
the  botany  of  his  time,  and  for  the  century  just  closed  full 
credit  should  not  be  withheld  from  those  whose  writings  fos- 
tered and  spread  an  interest  in  their  science.  Schleiden, 
Lindley,  Willkomm,  Gray,  Darwin,  Kerner  von  Marilaun, 
Gibson  and  Lubbock  have  shown  pre-eminent  ability  to  per- 
petuate the  old  and  awaken  new  interests.  Too  great  value 
can  scarcely  be  attributed  further  to  the  scientific  stimulus 
and  opportunity  due  to  the  publication  of  such  comprehensive 
class-books  as  the  general  text-book  of  Sachs,  the  Compara- 
tive Anatomy  of  DeBary,  the  physiological  manuals  of  Sachs 
and  Pfeffer,  the  pollination  works  of  Herman  Mueller  and 
the  dissemination  treatises  of  Haberland,  all  of  them  original 
contributions  to  science  as  well  as  adaptations  of  its  results  to 
the  purpose  of  the  teacher;  and  the  abridgments,  local 
adaptations,  popularizations  and  imitations  of  these  products 
of  leaders,  reaching  and  being  comprehended  by  a larger 
audience,  may  perhaps  have  done  even  more  toward  fanning 
into  flame  the  first  spark  of  enthusiasm  and  desire  for  re- 
search. 

Quite  as  noteworthy  is  the  advance  in  educational  and  in- 
structional methods,  and  appliances  other  than  books.  Up 
to  the  middle  of  the  century,  instruction  in  botany  was  con- 
fined to  more  or  less  perfunctory  lecture  courses,  and  the 
pupil  who  would  become  an  investigator  was  obliged  to  work 
out  his  own  salvation,  or  was  permitted  as  a special  favor  the 
privilege  of  association  with  a master.  The  opening  of  a 
botanical  laboratory  at  the  Univerity  of  Freiburg,  by  DeBary, 
in  1858,  marks  an  epoch.  It  is  a poor  college  to-day,  as  the 


140 


Trans . Acad.  Sci.  of  St.  Louis. 


equipment  of  colleges  now  goes,  which  has  not  a better  labor- 
atory and  a better  equipped  one  than  was  DeBary’s.  With 
the  introduction  of  laboratory  work  came  the  training,  in  the 
laboratories,  of  laboratory  teachers  to  spread  the  leaven,  not 
only  by  repeating  the  process  but  by  publishing  in  detail  their 
methods  for  the  benefit  of  others  who  could  not  work  under 
them.  It  would  be  impossible  to  overstate  our  debt  to  Huxley 
and  Martin’s  Biology  and  the  many  guides  of  which  it  was 
the  precursor,  to  Strasburger’s  Practicum,  the  various  treat- 
ises on  microscopic  technique  and  microchemistry,  and  the  in- 
creasing number  of  physiological  handbooks  which  have  grown 
out  of  Detmer’s  original.  That  the  botanical  world  has 
to-day  not  only  the  attainments  of  its  predecessors,  but  as  a 
regular  institution  these  facilities  which  did  not  formerly  ex- 
ist for  the  performance  of  work,  may  perhaps  be  regarded  as 
affording  ground  for  the  hope  that  the  century  upon  which 
we  have  now  entered  will  as  greatly  surpass  in  achievement 
the  one  just  closed  as  the  latter  did  all  of  its  predecessors. 

BOTANY  IN  THE  UNITED  STATES. 

Though  epitomized  in  the  preceding  general  survey  of  the 
field,  the  progress  in  our  country  of  what  has  been  called  the 
amiable  science  interests  us  so  directly  that  I may  briefly 
touch  on  it  in  conclusion. 

Systematic  phanerogamic  botany,  early  advanced  through 
the  labors  of  Nuttall,  Pursh,  the  Michaux,  Elliott  and  others, 
made  rapid  strides  about  the  middle  of  the  century, 
when  Torrey  and  Gray  undertook  the  publication  of  their 
Flora,  — unfortunately  never  completed,  partly  because 
of  the  wealth  of  new  material  brought  to  its  authors  as  a 
result  of  the  extensive  explorations  of  our  western  territory 
undertaken  by  the  Government.  Without  mentioning  others 
who  have  greatly  contributed  to  its  advancement  in  recent 
years,  I may  say  that  Gray’s  Manual,  Chapman’s  Flora  of 
the  Southern  States,  Watson’s  contributions  to  western  bot- 
any, Coulter’s  Kocky  Mountain  Botany,  and  the  masterly 
revisions  of  critical  groups  by  Gray,  Watson  and  Engelmann, 
have  brought  a knowledge  of  our  plants  within  the  reach 
alike  of  investigator  and  amateur ; while  few  countries  pos- 
sess a local  flora  comparable  with  that  of  Britton  and  Brown, 


Trelease  — Botany  During  the  19th  Century.  141 

and  the  great  Silva  of  Sargent,  now  nearly  completed,  stands 
quite  alone.  Eaton  and  Engelmann  laid  a good  foundation 
for  the  further  study  of  pteridophytes,  which  Davenport,  Rob- 
inson, Underwood  and  others  have  later  brought  to  the  hands 
of  every  working  botanist.  Through  the  work  of  Sullivant, 
Lesquereux,  James,  Austin,  Barnes  and  Underwood,  thebryo- 
phytes  have  been  similarly  put  within  easy  reach,  though  the 
current  work  of  Mrs.  Britton,  Evans,  Renauld  and  Cardot 
shows  that  even  more  than  with  the  superior  groups,  the  field 
for  systematic  research  is  here  still  open.  By  the  publica- 
tions of  Harvey,  Farlow,  Collins  and  others  on  marine  forms, 
and  of  Wood,  Wolle  and  others  on  those  of  fresh  water,  our 
algae  have  been  exceptionally  well  blocked  out.  Tucker- 
mann,  Willey  and  Williams  have  brought  the  lichens 
together:  and  though  less  advanced  than  either  of  the 

others,  the  great  group  of  fungi,  because  of  its  size, 
has  been  the  subject  of  more  actual  work  than  all  of  the 
remaining  cryptogams,  and  the  names  of  Berkeley,  an  Enlish- 
man,  and  of  Schweinitz,  Curtis,  Ravenel,  Farlow,  Thaxter, 
Peck  and  Ellis  stand  out  prominent  among  those  who  have 
contributed  to  its  lasting  literature.  Like  the  great  English 
botanists,  Americans  have  been  closer  adherents  to  the 
DeCandolle  classification  of  flowering  plants  than  to  the  later 
French  and  German  systems  until  very  recently;  but  the  dis- 
position of  to-day  is  strongly  toward  the  latter.  I may 
mention,  in  passing,  that  the  new  plantations  of  the  Missouri 
Botanical  Garden  will  be  twofold,  — one  portion  illustrating 
the  now  familiar  but  rapidly  passing  French-English  system, 
while  another  and  greater  part  will  follow  the  general  lines  of 
the  present  German  school. 

Americans  were  quick  to  take  up  the  Darwinian  ideas  of 
evolution,  — none  quicker  than  the  great  botanist  Asa  Gray, 
and  it  may  not  be  going  too  far  if  I say  that  nowhere  in  the 
world  has  horticultural  advantage  been  more  fully  taken  of 
their  teaching  than  in  America,  Bailey’s  varied  work  in  this 
field  being  particularly  mentionable. 

Though  morphological  teachings  were  prevalent  in  the  mid- 
dle part  of  the  century,  as  a research  subject  morphology  has 
been  confined  to  the  later  years,  during  which,  in  connection 


142 


Trans . Acad.  Sci.  of  St . Louis . 


with  more  precise  anatomical  studies,  it  has  contributed  to  an 
important  if  not  very  extensive  literature,  — largely,  it  must 
be  confessed,  resting  upon  the  studies  of  German-trained 
students. 

Vegetable  physiology,  as  a subject  for  serious  work  in  this 
country,  can  scarcely  be  traced  back  of  the  last  quarter  of  the 
century,  except  for  the  much  earlier  isolated  studies  of  Draper  ; 
but  to-day  the  force  of  several  well- equipped  laboratories, 
and  numerous  isolated  workers,  are  probing  the  difficult 
problems  the  solution  of  which  could  not  be  compassed  in  the 
century  just  closed.  Nowhere  has  that  phase  of  physiological 
work  known  as  bionomics  or  ecology  been  more  eagerly  taken 
up  than  in  this  country,  and,  beginning  with  Dr.  Gray,  a 
number  of  workers  have  enlarged  our  knowledge  of  the  pol- 
lination, dissemination  and  germination  of  plants,  while  the 
last  few  years  have  witnessed  a widespread  and  growing 
interest  in  the  vegetative  relations  of  plants  to  their  surround- 
ings, and  in  the  manner  in  which,  as  individuals  and  com- 
munities, they  compete  for  a foothold  on  the  earth. 

Without  going  into  details,  1 may  say  that  America  leads 
the  world  in  the  attention  given  to  botanical  (as  other)  re- 
search relating  to  agriculture  and  horticulture,  and  no  small 
part  of  the  recent  progress  in  this  field  has  come  from  our 
Government  and  State  laboratories  and  experiment  stations. 

In  conclusion,  as,  perhaps,  the  greatest  advance  in  botany 
made  in  this  country  during  the  century,  I may  note  the  in- 
crease and  improvement  in  means  and  methods  for  instruction 
The  great  strides  made  in  this  direction  by  the  Germans  at 
the  close  of  the  Franco-Prussian  war,  and  the  prestige  of 
DeBary,  Sachs,  Pfefter  and  Strasburger  in  their  Universities, 
stimulated  and  attracted  Americans  to  such  an  extent  that 
to-day  no  country,  aside  from  Germany,  offers  so  many,  so 
good,  or  so  varied  opportunities  for  training  in  scientific  bot- 
any as  we  possess  in  the  United  States,  and  a rich  fruition 
may  be  confidently  expected  in  the  century  on  which  we  have 
now  entered. 


Issued  November  26,  19  01. 


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